Effect of microbial films on settlement of bryozoan larvae (Bugula simplex,B. stolonifera and B. turrita)

The influence of microbial films on larval settlement was studied for sympatric species of cellularioid cheilostome bryozoans (Bugula simplex, B. stolonifera, and B. turrita). Settlement experiments were conducted in the laboratory in 1981 with and without a choice of filmed or unfilmed substrates. When given a choice of substrates, the larvae of all three bryozoan species significantly preferred the filmed substrate. When not given a choice of substrates, both B. simplex and B. turrita larvae tended not to settle on the unfilmed substrate, while the filmed and unfilmed substrates were equally attractive to B. stolonifera. Microbial films enhanced the settlement of all three species.     

Ancient behaviors of larval amphibians in response to an emerging fungal pathogen, <Emphasis Type="Italic">Batrachochytrium dendrobatidis</Emphasis>

Behaviors have evolved in response to various selection pressures over evolutionary time. However, not all behaviors are adaptive. Some presumably “ancient” behaviors, persistent for millions of years, may be detrimental in the face of novel selection pressures in modern times. These pressures include a multitude of emerging infectious diseases which may be stimulated by environmental changes. We examined how a globally emerging amphibian pathogen, Batrachochytrium dendrobatidis (BD), affected two key evolutionarily persistent behaviors displayed by amphibian larvae: aggregation and thermoregulation. Larval aggregation behavior is often essential for foraging, thermoregulation, and antipredator defense, but varies among species. Thermoregulatory behavior speeds larval development in ephemeral habitats. Specifically, we examined whether aggregation and thermoregulatory behaviors changed when exposed to the BD pathogen in two species (Bufo boreas and Rana cascadae) whose larvae aggregate in nature. In laboratory choice tests, larvae of neither species avoided infected conspecifics. BD-exposed B. boreas larvae aggregated, while unexposed R. cascadae larvae associated more frequently with BD-exposed conspecifics. There was no evidence of behavioral fever or altered thermoregulation in larvae of four species we examined (Pseudacris regilla, Rana aurora, B. boreas, R. cascadae). The absence of behavioral fever may suggest an inability of the larvae of some host species to mediate infection risk by this pathogen. Thermoregulatory behaviors may exhibit a high degree of evolutionary inertia in amphibian hosts because they are linked with host physiology and developmental rates, while altered aggregation behaviors could potentially elevate pathogen transmission rates, leading to increased infection risk in social amphibian species.     

Effect of bryozoan colonization on inorganic nitrogen acquisition by the kelps Agarum fimbriatum and Macrocystis integrifolia

The effect of bryozoan colonization on inorganic nitrogen acquisition by Agarum fimbriatum Harv. and Macrocystis integrifolia Bory., collected from the west coast of Vancouver Island, British Columbia, Canada, was examined in laboratory experiments during June and July 1992. Pieces of kelp blades that were completely covered on one side by the bryozoans Lichenopora novae-zelandiae Busk or Membranipora membranacea, L., or uncolonized (clean treatment), were used to estimate the rate at which nitrate and ammonium were removed from the surrounding seawater. In addition, the rate of ammonium excretion by bryozoans isolated from their associated kelp was measured and also estimated from the results of the uptake experiments. Values obtained were used to estimate the contribution of ammonium excreted by bryozoans to the total amount of inorganic nitrogen available to the associated kelp. Both bryozoan species reduced the ability of the associated kelp to remove nitrate and ammonium from seawater but provided a source of ammonium to the kelp through excretion. The nitrogen status of colonized and clean kelp disks was determined from the ratio of total particulate carbon to total particulate nitrogen (C:N ratio). The C:N ratios for A. fimbriatum colonized with either L. novae-zelandiae or M. membranacea were similar (C:N=12 to 14), and differences between colonized and clean treatments were not significant. For A. fimbriatum, therefore, the C:N ratio indicates that this species was not nitrogen limited at the time of the present study. In contrast, both colonized and clean disks of M. integrifolia were nitrogen limited, but colonized disks (C:N=19) were significantly less limited by nitrogen than clean disks (C:N=29). Results are discussed in relation to the different environments inhabited by both kelp species and are consistent with the hypothesis that ammonium excreted by bryozoans was an important source of inorganic nitrogen to M. integrifolia, but not to A. fimbriatum, at the time of the study.     

Inhibition of attachment of larval barnacles,Balanus amphitrite,by bacterial surface films

Films of bacteria on solid substrata can positively or negatively influence the attachment of marine invertebrate larvae. Effects of marine bacteria on the attachment of cypris larvae of the barnacle Balanus amphitrite Darwin were examined in the laboratory. Bacteria, grown to mid-exponential phase and allowed to adsorb irreversibly to polystyrene petri dishes, attached in densities of 10⁷ cells cm^-2. Assays (22h) were used to compare the effects of adsorbed cells of 18 different bacterial species on larval barnacle attachment. Most of the adsorbed bacteria either inhibited or had no effect on larval attachment compared to clean surfaces. Experiments testing the effect of larval age on barnacle attachment were conducted with six species of bacteria and showed that older larvae attached in higher percentages to clean surfaces and that bacterial films generally inhibited larval attaschment. Both the species of bacteria and the in situ age of the adsorbed bacteria affected barnacle attachment response: older films of Deleya (Pseudomonas) marina were more inhibitory. Bacterial extracellular materials may be involved in the inhibitory process.     

Larval recruitment in response to manipulated field flows

Settlement responses to boundary-layer flow of several invertebrate taxa, including the hydroid Tubularia crocea, the bryozoans Bugula turrita and Schizoporella unicornis, and the tube-building polychaete Hydroides dianthus were studied in manipulated field flows in Great Harbor, Massachusetts, USA. During three experiments in 1989 and two in 1990, densities of newly-recruited larvae were measured on flat plates, whose flow regimes had been manipulated by altering the leading-edge configurations. Settlement responses to flow were strongly species-specific, with T. crocea preferring regions of both high turbulence and strong shear stress, and S. unicornis settling a most exclusively in regions of high shear stress. B. turrita settled most prominently in regions of reduced shear stress, exhibiting settlement patterns that closely approximated predictions from a model of passive particle contact. H. dianthus showed a moderate avoidance of regions with high shear stress. These results indicate that boundary-layer flows affect settlement of several common encrusting species, a probable consequence of larval behaviors such as substrate rejection or exploration in response to flow. These responses are likely to generate patchiness during initial colonization of natural habitats, and certainly affect colonization of settlement panels commonly used in marine ecological studies.     

Reproductive strategies and life histories in the cheilostome marine bryozoans Chartella papyracea and Bugula flabellata

Seasonal life history phenomena were monitored through 1978 for two sublittoral cheilostome bryozoans from souther Britain: Chartella papyracea (Ellis and Solander), a non-placental brooder, and Bugula flabellata (Thompson in Gray), which is placental. The intracolonial relationships between growth by zooid budding, polypide recycling, and sexual reproduction, were analysed for both species — quantitatively in the case of C. papyracea. Although both species grow only in summer, their life histories differ. C. papyracea is more of a K-strategist, producing perennial fronds that release larvae in several successive years, most being liberated in winter months. The reduced need for maximization of the larval productivity rate is reflected in the intracolonial configuration of polypide and sexual recycling. B. flabellata is more of an r-strategist. The larvae are produced by the ephemeral fronds of two successive colony generations each summer. Colonies are dormant in winter. The need to maximize the larval productivity rate in this species is reflected in the polypide/sexual recycling configuration. It is believed that embryonic placentation contributes to the maximization of the larval productivity rate in this and other placental brooders.     

Origin,composition, and function of secretions from pyriform organs and internal sacs of four settling cheilo-ctenostome bryozoan larvae

Settling larvae of the bryozoans Flustrellida hispida, Flustra foliacea, Alcyonidium polycum, and Bugula neritina exhibit a common behavioral pattern. A settlement site is selected after initial exploration, and temporary attachment is accomplished with a sticky acid mucopolysaccharide secretion exuded from the pyriform organ. Permanent attachment is accomplished by acid mucopolysaccharide and protein secretions of the everted internal sac.     

Distribution patterns of larval myctophid fishes in the transition region of the western North Pacific

We examined the species composition and horizontal distribution of myctophid fish larvae in the transition region of the western North Pacific during the early summer. In total, 4,760 myctophid larvae were collected at 44 stations; 18 species of myctophids from 15 genera were collected, and the 8 most abundant species accounted for >95% of larvae. The distribution patterns of these larvae were well defined by the hydrographic structures of the study area including the Oyashio and Kuroshio fronts, the Subarctic Boundary, and a warm core ring. The horizontal distribution patterns of the eight dominant species were categorized into three groups: northern transition water (Stenobrachius nannochir, Tarletonbeania taylori, and Lampanyctus jordani), southern transition water (Symbolophorus californiensis, Diaphus theta, and Nannobrachium regale), and Kuroshio (Myctophum asperum and Diaphus garmani). The Subarctic Boundary defined the distributions of the northern and southern transition-water groups. The importance of areas of western North Pacific transition water as spawning and nursery grounds for subarctic, transitional, and subtropical myctophid fishes was indicated by the relationship between the horizontal distribution patterns of larvae, juveniles, and adults and the physical oceanographic structures.Communicated by T. Ikeda, Hakodate     

Onset of symbiosis and distribution patterns of symbiotic dinoflagellates in the larvae of scleractinian corals

The establishment of symbiosis in early developmental stages is important for reef-building corals because of the need for photosynthetically derived nutrition. Corals spawn eggs and sperm, or brood planula larvae and shed them into the water. Some coral eggs or planulae directly inherit symbiotic dinoflagellates (Symbiodinium spp.) from their parents, while others acquire them at each generation. In most species examined to date, the larvae without dinoflagellates (aposymbiotic larvae) can acquire symbionts during the larval stage, but little is known regarding the timing and detailed process of the onset of symbiosis. We examined larval uptake of symbiotic dinoflagellates in nine species of scleractinian corals, the onset of symbiosis through the early larval stages, and the distribution pattern of symbionts within the larval host, while living and with histology, of two acroporid corals under laboratory conditions. The larvae acquired symbiotic dinoflagellates during the planktonic phase in all corals examined which included Acropora digitifera, A. florida, A. intermedia, A. tenuis, Isopora palifera, Favia pallida, F. lizardensis, Pseudosiderastrea tayamai, and Ctenactis echinata. The larvae of A. digitifera and A. tenuis first acquired symbionts 6 and 5 days after fertilization, respectively. In A. digitifera larvae, this coincided with the formation of an oral pore and coelenteron. The number of symbiotic dinoflagellates increased over the experimental periods in both species. To test the hypothesis that nutrients promotes symbiotic uptake, the number of incorporated dinoflagellates was compared in the presence and absence of homogenized Artemia sp. A likelihood ratio test assuming a log-linear model indicated that Artemia sp. had a significantly positive effect on symbiont acquisition. These results suggest that the acquisition of symbiotic dinoflagellates during larval stages is in common with many coral species, and that the development of both a mouth and coelenteron play important roles in symbiont acquisition.     

A study on the effects of four heavy metals on five species of chironomous larvae present on Mahé Island of Seychelles

A study was conducted on the island of Mahé, Seychelles, on five species of chironomous larvae endemic to the island. The chironomous larvae were exposed to four heavy metals – copper (Cu), cadmium (Cd), mercury (Hg), and zinc (Zn), for a 96-hr period, and the LC₅₀ values were calculated. It was observed that the most sensitive species of chironomous larvae was Gymnometriocnemus mahensis. The least sensitive form of chironomous larvae examined was Paramerina minima. The 96-hr LC₅₀ values obtained for Tanypus complanatus for Cu, Cd, Hg, and Zn were 1.9 (1.73–2.09), 1.35 (1.17–1.57), 0.33 (0.26–0.43), and 28.96 (27.03–31.03) ppm, respectively. Similar results were obtained for other species studied.     

Behaviour of echinoid larvae around sharp haloclines: effects of the salinity gradient and dietary conditioning

We examined larval response to a range of sharp haloclines and determined the effect of dietary conditioning on that response in the sea urchins Echinometra lucunter and Arbacia punctulata. We reared larvae in the laboratory under a high or low concentration of either single (Isochrysis galbana) or mixed (Isochrysis galbana, Dunaliella tertiolecta, Thalassiosira weissflogii) microalgal species. For both species of sea urchins, rate of larval development was faster and age-specific larval length and width were greater in high-ration than low-ration diets. We examined the distribution of two- and four-arm larvae of E. lucunter from each diet treatment and of four-arm larvae of A. punctulata from the high-ration diets in cylinders with experimentally constructed haloclines. In three of the halocline treatments, the salinity of the bottom layer was 33‰ and that of the top layer was 21, 24 or 27‰ (21/33, 24/33 and 27/33) and in a fourth one, the salinities of the bottom and top layer were 30 and 21‰, respectively (21/30). The position of larvae in the cylinders varied with the steepness of the halocline and with dietary conditioning for both sea urchin species and all developmental stages tested. Significantly more larvae crossed the haloclines into water of 24 and 27‰ salinity than into water of 21‰ salinity. We observed an effect of diet on the position of larvae in the cylinders, and that effect varied among halocline treatments for both species. The proportion of larvae of E.lucunter that crossed the halocline was greater in low- than high-ration diets in the 24/33 and 27/33 treatments. Position of four-arm larvae in the cylinders also varied with food quality in high-ration diets: for E.lucunter in the 24/33 treatments, and for A. punctulata in the 21/30 treatments, more larvae from the single- than from the mixed-species diets were present above the halocline. Salinity in the adult habitat during most of the active reproductive period ranged from 15 to 40‰. We showed that larvae can respond to gradients in salinity, and therefore can remain within a water mass of higher salinity overlying the adult habitat. However, survival of poorly fed larvae may be increased if they are introduced into a new water mass and carried away from a nutritionally poor environment. Received: 9 July 1997 / Accepted: 12 January 1998     

Inception of formation and early morphogenesis of the bacterial light organ of the sea urchin cardinalfish, <Emphasis Type="Italic">Siphamia versicolor</Emphasis>

The cardinalfish Siphamia versicolor (Perciformes: Apogonidae) forms a bioluminescent symbiosis with the marine luminous bacterium Photobacterium mandapamensis, harboring the bacteria in a ventral, disc-shaped light organ and using the bacterial light apparently for counterillumination and attracting prey. Little definitive information has been available on the developmental and microbiological events surrounding the initiation of symbiosis, a critical stage in the life history of the fish, in S. versicolor or any of the many other species of bacterially luminous fish. To identify the stage at which light organ formation begins, to determine the origin of cells forming the light organ, and to characterize its bacterial colonization status during development, early developmental stages of S. versicolor obtained and reared from wild-caught mouth-brooding males were examined with histological and microbiological methods. A light organ primordium was not evident in embryos, post-embryos, or pre-release larvae, whereas the light organ began to form within 1 day of release of full-term pre-flexion larvae from the mouths of male fish. Analysis of post-release larvae revealed that the light organ arises from a proliferation and differentiation of intestinal epithelial cells, and that it quickly develops structural complexity, including the formation of chambers and gaps contiguous with the intestinal epithelium. However, the nascent light organ remained uncolonized by the symbiotic bacteria through several days of post-release development, even in the presence of high numbers of the symbiotic bacteria. These results demonstrate that the inception of light organ formation in S. versicolor occurs independently of its symbiotic bacteria and that receptivity to bacterial colonization apparently requires substantial post-release development of the light organ. Larvae therefore most likely acquire their symbiotic bacteria from seawater, during or shortly after the transition from the pre-flexion to the flexion developmental stage.     

Reproductive cycles and larval morphology of three Recent species of Argyrotheca (Terebratellacea: Brachiopoda) from Mediterranean submarine caves

Although the three examined species of Argyrotheca are quite common in the Mediterranean, little is known about their reproductive cycles. This study shows that Argyrotheca cordata and Argyrotheca cistellula have continuous breeding activity, indicated by the occurrence of ripe eggs and larvae in their brood pouches throughout the year. The absolute number of larvae found in the brood pouches of both species corresponds to the adult's size. The third species examined, Argyrotheca cuneata, tends to breed in autumn, with larvae only present in September and November, but the sample-size was small. Larval development is very similar in all three Argyrotheca species, each embryo going through a gastrula stage, a two-lobed and a three-lobed stage. Before leaving the brood pouch, the larvae have an apical lobe with a girdle of elongated cilia, a mantle lobe with a midventral band of cilia and a pedicle lobe without ciliation. The larvae of A. cordata and A. cuneata possess four bundles of larval setae, whereas A. cistellula shows no setae at all. Received: 6 November 1998 / Accepted: 15 March 1999     

Substratum preferences in planula larvae of two species of scleractinian corals, Goniastrea retiformis and Stylaraea punctata

To test whether coral planulae recruit randomly to different coral reef habitats or have specific substratum preferences, the settling behavior of planulae from two shallow water coral species from Pago Bay, Guam (13°25.02N, 144°47.30E) were examined in the laboratory in June and July of 1995. Goniastrea retiformis is generally restricted to the shallow reef front (<10 m depth) in areas dominated by crustose coralline algae (CCA), while Stylaraea punctata is abundant on inner reef flats were CCA coverage is low and sand and carbonate rubble covered by biofilms is common. When presented with four substrata (1) carbonate rock scrubbed free of biofilm and dried as a control, (2) the CCA Hydrolithon reinboldii, (3) the CCA Peyssonelia sp., and (4) naturally conditioned carbonate rubble covered by a biofilm, G. retiformis larvae showed a significant preference for H. reinboldii, and S. punctata larvae for the carbonate biofilm treatment. The preference shown by S. punctata larvae for biofilmed surfaces did not diminish with increasing larval age up to 11 days. These results suggest that the larvae of both species are capable of habitat selection, and that the preferred substrata among those tested bears a relationship to the habitats in which adult colonies were found. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effect of the receding ice-edge on the condition of mid-water fishes in the northwestern Weddell Sea: results from biochemical assays with notes on diet

Electrona antarctica and Bathylagus antarcticus are dominant fishes in the Southern Ocean pelagic ecosystem with disparate life histories, making them excellent subjects for comparative studies. Twenty-one indices of nutritional condition were compared in both species by observing changes in each index as it varied spatially and temporally in association with the marginal ice-zone bloom in the northwestern Weddell Sea. E. antarctica and B. antarcticus exhibited very different patterns of response to the bloom. Nutritional condition increased in 11 of 21 measures in E. antarctica, suggesting that it was in poorer condition at ice-covered stations and in greatly improved condition in post-bloom, open-water stations. The data, combined with a few observations of larvae, indicate that E. antarctica increases feeding before the spring bloom, rapidly building stores of lipid, which are probably used for spawning. In contrast, only three measures of condition were variable in B. antarcticus, and lipid actually decreased from ice-covered to open water. RNA:DNA values increased in open water, coinciding with an increase in food volume present in their guts. These observations, coupled with observations of larvae in ice-covered stations, suggest that B. antarcticus possesses sufficient energetic stores to spawn prior to the bloom, so that larvae are able to fully utilize available resources associated with the spring bloom. Increased somatic growth, as indicated by RNA:DNA, may not begin until later, when the bloom was beginning to decline. Earlier work, which suggested that the deeper living, non-migrating species, like B. antarcticus, would not be affected by the bloom until after shallower, migratory species, like E. antarctica, is supported. Of the assays tested, RNA:DNA appeared to be the most sensitive. The combined assessment of many measures including biochemical, compositional, dietary, and age estimates from single specimens is possible if the samples are treated with sufficient care. The reliance upon a single measure to help interpret the ecology of a species, especially in nekton-sized species, is not as effective as techniques used in combination. Received: 12 January 2000 / Accepted: 16 August 2000     

Comparative fertilization biology of gymnolaemate bryozoans

Locations and oogenic stages of eggs at the time of sperm-egg fusion were determined for nine gymnolaemate species exhibiting different reproductive strategies with respect to site of embryonic development (e.g., water column, introvert, embryo sac, or ovicell) larval type, and zooid sexuality. Ovarian, coelomic, and recently spawned oocytes obtained from freshly collected colonies between 1988 and 1995 were stained with aceto-orcein and examined for the presence of a sperm nucleus. All nine species are shown to have a similar fertilization biology in which sperm fuse with eggs in or near ovaries and egg activation is delayed until eggs are spawned. In species such as the etenostomeAlcyonidium sp. and the cheilostomeElectra pilosa (Linnaeus, 1767), which produce marierons oocytes that are spawned into the water column and have cyphonautes larvae, sperm fuse with late growth stage ovarian oocytes after germinal vesicle breakdown (GVBD) at or near ovulation. In species that produce few oocytes, brood embryos, and release coronate larvae, sperm fuse with late growth stage ovarian primary oocytes before GVBD, as in the ctenostomeBowerbankia gracilis Leidy, 1855, or with very early growth stage ovarian primary oocytes, as in the cheilostomesCribrilina corbicula (O'Donoghue, 1923),Dendrobeania lichenoides (Robertson, 1900),Hippodiplosia insculpta (Hincks, 1882),Schizoporella serialis (Heller, 1867),T ricellaria gracilis (Smitt, 1867), andWatersipora arcuata Banta, 1969. Fertilization success was high in all nine species, with 83 to 100% of oocytes at an appropriate oogenic stage containing a sperm nucleus or early male pronucleus. Gymnolaemate bryozoans may maintain high levels of fertilization success by: (1) concentrating spawned sperm from the water column using colony feeding currents; (2) localizing sperm-egg encounters in or near the ovary, (3) using sperm when available by permitting sperm to fuse with early growth stage oocytes; and (4) selfing.     

Clearance rates of food suspension and food passage rates as a function of temperature in two north-sea bryozoans

The filtration rate has been determined for Electra pilosa and Conopeum reticulum, acclimated at 4 constant temperatures (6°, 12°, 18°, 22°C). Cryptomonas sp. was used as the test food-organism. Temperature affects filtration rate in both bryozoans. Clearance rate, measured at hourly intervals during the course of the experiment, was never uniform. Possibly, temporary reductions in ciliary activities were induced by the artificial conditions under which the experiments were carried out. The results obtained compare well with those reported by Bullivant (1968) on Bowerbankia imbricata, a cosmopolitan ctenostome. Rate of food passage through the alimentary canal is affected by temperature. The present study is the first to demonstrate the effect of temperature on the rate of filtration in bryozoans.     

Population genetic consequences of larval dispersal mode and hydrography: a case study with bryozoans

Genetic analysis of the marine bryozoans Celleporella hyalina and Electra pilosa using the RAPD technique revealed population structuring corresponding to the contrasting modes of larval dispersal. Samples of C. hyalina exhibited genetic differentiation over distances as small as 10 m, concordant with the limited dispersal predicted by a simulation model, based on the short pelagic phase of the lecithotrophic larvae and the local hydrography. In contrast, E. pilosa showed high levels of genetic heterogeneity only over much larger spatial scales, commensurate with its production of comparatively long-lived planktotrophic larvae. The population differentiation observed between samples of E. pilosa, collected from sites 70 km apart, is reconcilable with coastal water currents and frontal systems that restrict the exchange of water masses between the two sites. Hydrographic conditions and discontinuous distribution of suitable substrata therefore are seen to constrain gene flow, creating opportunities for local genetic differentiation despite the high dispersal potential of pelagic larvae. Received: 9 August 2000 / Accepted: 18 November 2000     

Chemoecological studies of the exocrine glandular larval secretions of two chrysomelid species (Coleoptera):Phaedon cochleariae andChrysomela lapponica

Summary The exocrine glandular secretions of larvae of the subfamily Chrysomelinae are known to repel conspecific adults, other competitive phytophagous insects and natural enemies. InPhaedon cochleariae, the intraspecific activity of tlc fractions of the larval secretion was tested in order to examine the ecological significance of two fractions containing minor components and a fraction containing the major compound, the cyclopentanoid monoterpene (epi)chrysomelidial. InChrysomela lapponica, the defensive activity of the larval secretion against ants is known from specimens feeding upon willow or birch. The feeding preferences of larvae and adults ofC. lapponica from a Finnish and a Czech population were tested. The Finnish individuals significantly preferred feeding uponSalix borealis, whereas they hardly fed upon birch. The Czech specimens clearly preferred birch (Betula pubescens) to willow species. Application of salicin onto leaves of a willow species free of this phenolglycoside revealed that the Finnish individuals preferred feeding upon leaves with salicin. On the other hand, the Czech individuals avoided feeding upon leaves ofB. pubescens treated with salicin. The chemical composition of the glandular secretion of the Finnish larvae differed from the one of the Czech larvae. GC-MS-analyses of the secretions revealed that salicylaldehyde was the only major component of the secretion of Finnish larvae feeding upon the salicin-containing willowS. borealis. The glandular secretion of the Czech larvae feeding upon birch contained numerous esters of isobutyric acid and 2-methylbutyric acid. When Czech larvae had fed upon a salicin-containing willow (S. fragilis), the major compounds of their secretion were benzoic acid, salicylalcohol and benzoic acid esters; salicylaldehyde was only detected in traces. Thus,C. lapponica individuals from the Finland population adapted so closely to a salicincontaining willow that they clearly prefer this plant for food and that they obviously derive their main larval defensive compound (salicylaldehyde) from their host-plant.